Early Natural History of the Greater Glider, Petauroides volans (Kerr, 1792) K. Shane Maloney' and Jamie M. Harris^ NSW ' School ofBiological Sciences, University ofWollongong, 2522 ([email protected]); NSW ^School ofEnvironmental Science and Management, Southern Cross University, Lismore 2480 ([email protected]) Maloney,K.S. andHarris, J.M. (2008). Earlynaturalhistoryofthegreaterglider,Petauroidesvolans(Kerr, 1792).Proceedings oftheLinnean SocietyofNewSouth Wales 129, 39-55. Early accounts ofthe greater glider Petauroides volans (Marsupialia; Pseudocheiridae) are reviewed, startingwithArthurPhillips' 1789accountin The VoyageofGovernorPhilliptoBotanyBayandproceeding to the latest taxonomic works. This species has a quite complicated and confusing taxonomic history. It has been listed as a member ofno fewer than 1 genera with about 23 different binomial names since its discovery. Inthispaper,wereviewsomeofthistaxonomiccomplexityandearlydescriptionsofthespecies' morphology, dentition, behaviour, distribution andabundance. We foundthattaxonomic descriptions oiP. volans have been frequently confused with those ofa number ofother gliding possums, particularly the yellow-belliedgWdsrPetaurusaustralis. EarlydescriptionsofthemorphologyofP. volansweregivenonly inbroadgeneralterms. Morevalue canbeplacedontheearlybehavioural observations, andonthe earliest records ofits occurrence. This paper examines some ofthe oldest accounts ofP. volans and assesses their significance. Manuscriptreceived 14 Februry2007, acceptedforpublication 12 December2007. KEYWORDS: natural history, nomenclature, Petauroides volans, Pseudocheiridae. included in many ofthe earliest zoological accounts INTRODUCTION of the Australian fauna. However, few modem zoologists are aware of the historical significance The greater glider, Petauroides volans and value ofthis old literature as it relates to this and (Marsupialia: Pseudocheiridae), is the largest gliding other species (see also Harris 2006). Whilst some marsupial and is endemic to eastern mainland of this literature on P. volans has been reviewed by Australia (McKay 1995). Currently, there are two McKay (1982), this was limited to aspects of the recognised sub-species: P. volans volans, which nomenclature of the genus name Petauroides (and occurs in south eastern Australia (from Victoria in also Petaurus). In this contribution, we have sought the south, through mainly coastal New South Wales to provide a comprehensive survey of the early (NSW) to the Rockhampton district in north-east natural history literature pertaining to P. volans, Queensland (Qld)); and P. v. minor, which occurs in including information on discovery, taxonomy, very farnorth-eastQld(fromthe DawsonRivertothe dentition, morphology, distribution, abundance, diet BarronRiver) (Flannery 1994). Itis aroundthe size of andbehaviour. a domestic cat, with females being larger than males (Flannery 1994; Kavanagh and Wheeler 2004). Most TAXONOMYAND NOMENCLATURE individuals are jet black on the dorsum and creamy white on the ventrum, but pure white forms are not uncommon and intermediate colours are also found Governor Arthur Phillip reported 'black flying (Flannery 1994; McKay 1995; Lindenmayer 2002). opossum'fromNSW(Phillip 1789).Amale specimen This species is nocturnal, arboreal and folivorous owned by Henry Constantine Nowell was illustrated and is dependent on tree hollows for its nesting (Figure 1), although no details on the precise requirements. collection locality were published. Presumably it was Petauroides volans is relatively conspicuous and found in the vicinity of Port Jackson. Phillip (1789, was quickly noticed by the early colonists (Phillip 1790) recognised that it represented a new species 1789). Subsequently, descriptions ofthis specieswere and suggested taxonomic affinity with American EARLY HISTORY OF THE GREATER GLIDER Figure 1: Blackflyingopossum {=Petauroides volans) drawn by P. Mazellandpublished in Phillip (1789). Note the opposable clawless hallux and syndactylous digits on each ofthe hind feet. 40 Proc. Linn. Soc. N.S.W., 129, 2008 MALONEYAND HARRIS K.S. J.M. Didelphis, although a specific name was not offered. proposed the name D. macronra. In Shaw's work. A few years later, Kerr (1792) named Phillip's The Zoology ofNew Holland, a juvenile specimen specimen Didelphis volans, derived from the Latin drawn by James Sowerby was illustrated (Figure 2). word 'volare' meaning 'to fly' (Strahan 1981). A Shaw (1800) explained that it was sent to him by year later, Meyer (1793) named Phillip's specimen John White, who was the first Surgeon-General for D. voluccella, and a year later still, Shaw (1794) the colony ofNSW. Figure 2: Long-tailed opossum Didelphis macroum {=Petauroides volans) from Zoology ofNew Holland (1794) by George Shaw. The figure was drawn by James Sowerby. This illustration was also reproduced in Shaw (1800) and Desmarest (1820). Proc. Linn. Soc. N.S.W., 129, 2008 41 EARLY HISTORY OF THE GREATER GLIDER Cuvier (1798) followed use of the name D. directly, and obtained a photograph ofthe specimen volans (Kerr 1792) but questioned the affiliation (number CGI990-408) and although no patagium withthe genusDidelphis. Nevertheless, Shaw (1800) was evident in the photograph, it looks like a greater continued the use ofZ). macroura. Bechstein (1800) glider because ofits substantially long tail and hairy elevated the name Voluccella, usedby Meyer (1793), ears. The arrangement of Desmarest's (1817) was to generic level and proposed V. nigra for the subject later followedby Cuvier (1826), Lesson (1827, 1828, species, but he evidently confused the greater glider 1830, 1838), and Fischer (1829). Bennett (1837) also and the yellow-bellied glider Petaurus australis in used Desmarest's (1817) terminology, although he synonymy. His proposed V. nigra incorporated D. appears to have used P. peronii in reference to the voluccella Meyer, 1793 (=Petauroides volans) and sugar gliderPetaurus breviceps. "Hepoona Roo" White, 1790 {=Petaurus australis). Desmarest(1820)appliedPetauristatosupercede It is understood that Hepoona Roo is P. australis Petaurus, and maintained Petaurista taguanoides, P. and not Petauroides volans (McKay 1982, 1988). macroura and P. peronii as separate species (later Bechstein (1800) also advanced V. macroura as a followed by Cuvier 1827, 1829). However, this was separate species that incorporated D. volans Kerr, flawed as Petaurista had been advanced for the giant 1792 and D. macroura Shaw, 1794. Thus, V. nigra flying squirrels (Rodentia) by Link (1795) (see also and V. macroura are both synonyms of P. volans. Fischer 1814; Thomas 1888; Sherbom 1902; Palmer Voluccella Bechstein, 1800 was discontinued for the 1904). Waterhouse (1838b), Gloger (1842), Gould subject species because this genus name had already (1863) andThomas (1885), persistedwiththis invalid beenadvancedbyFabricius (1794) fora species offly generic name for the greater glider. (Diptera: Bombyliidae) (Thomas 1888; McKay 1988; Frederic Cuvier (1825) mentioned Petaurus Evenhuis 1991). Hence, Voluccella Bechstein, 1800 didelphoides Geoffroy, an apparent new name forthe is ajunior generic synonym for Petauroides but not subject species (Thomas 1888; Iredale andTroughton Voluccella Ydibncm?,, 1794. 1934; de Beaufort 1966). However, later works by Phalanger volans was used by Lacepede (1801), F. Cuvier and also his brother Georges, made no whilst Desmarest (1803) and Tiedemarm (1808) reference to P. didelphoides (Cuvier 1826, 1827, placed it under Phalangista (see also Schinz 1821; 1829). de Beaufort (1966) noted that Cuvier (1825) Thomas 1888). Turton (1806) mistakenly thought offered no specific descriptions, and stated that he that the descriptions by Kerr (1792: D. volans) and was unable to find any reference to Geoffroy as the Shaw (1794: D. macroura) represented two separate authority forthe name. It is uncertain whether Cuvier species. Oken (1816) made a similar mistake, but intended this name for the greater glider. Iredale and also erroneously included Petaurus australis in the Troughton (1934) considered it a vernacularname. synonymy for one ofhis proposed species. This was Lesson (1828, 1830, 1838) listed the "Black Petaurus niger, and the epithet was a gender change Flying Opossum" of Phillip (1789) (=the greater of Bechstein's (1800) nigra (see also Iredale and glider) as ajunior synonym ofPetaurus taguanoides. Troughton 1934). Oken's (1816) second species was This was subsequently repeated by Fischer (1829), Petaurus macroura. Wagner (1843), Schinz (1844) and Giebel (1859). Desmarest (1817) listed three species {Petaurus Waterhouse (1838a)thenstatedthattwo specimens of macrourus, P. peronii and P. taguanoides). Petaurus Petaurista taguanoides were held in the Museum of macrourus included a slight change in the epithet to theZoologicalSocietyofLondon(ZSL), oneofwhich standardise the gender ofthe binomial. Desmarest's was a 'white variety'. Waterhouse's (1841) included explanationthatthemembraneofP.pero/?//"terminates an illustration ofa greater glider(Figure 3) and stated at the elbow" is good evidence that this specimen that "Specimens which are totally white, and others was also P. volans. For P. taguanoides however, the which are white and irregularly variegated with grey, synonymy was confused with the yellow-bellied are not rare". Waterhouse (1841) was wrong when Glider [i.e. Didelphis petaurus of Shaw (1791) and he suggested that P. macrourus is P.flaviventer {=P. "HepoonaRoo"ofWhite (1790)] andthe descriptions australis) (see alsoWagner 1855; Giebel 1859; Gould aboutthepatagiumendingatthewristsuggestedtous 1863).Descriptionsoftaguanoidesspecimensinmany thatthisspecimenwasnotthegreaterglider.However, 19* century publications subsequent to Waterhouse accordingto thepublications ofthe MuseumNational (1841) appear to represent the greater glider (e.g. d'HistoireNaturelle (MNHN) thetype specimenofP. Owen 1841, 1845; Gloger 1842; Waterhouse 1846; taguanoides as described by Desmarest is indeed P. Gervais 1855; Gerrard 1862; Brehms 1880; Flower volans (de Beaufort 1966; Julien-Laferriere 1994). To 1884; Forbes-Leith and Lucas 1884; Krefft 1864; MNHN confirm this identification we contacted the Haswell 1886; Jentink 1886; Lucas 1890). 42 Proc. Linn. Soc. N.S.W., 129, 2008 . K.S. MALONEYAND J.M. HARRIS 1888; Iredale and Troughton 1934; McKay 1982). However, McKay (1988) stated that P. leucogaster was Tncertae sedis' (of uncertain position) because the specimen could no longer be found at the AM. He suggested that the locality for Mitchell's specimen was outside the range ofP. volans and may have been Petaurus norfolcensis M.R. Oldfield Thomas, of the British Museum of Natural History (BMNH), revised the taxonomy of the subject species several times duringtheperiod 1879-1923.Thomas (1879) noted that the specific name volans Kerr antedated taguanoides Desmarest, and maintained that the correct binomial was Petaurus A volans. fewyearslater,however,he listeditasPetauristavolans(Thomas 1885). After finding that Petaurista was unavailable, Thomas (1888) advanced Petauroides to replace the previous generic names. He listed two subspecies: Petauroides volans typicusasthe southernform; andi^ v. minorasthenorthernform(following Collett 1887). Later, Thomas (1923) received further examples from Qld and considered that there were two additional subspecies: P. v. incanus andP. V. armillatus. Thomas (1923) mentioned that Ogilby (1892) referred to "Dr Ramsay's P. cinereus" and that it "seemsnevertohavebeendescribed". However, Ramsay (1890) did indeed Figure3:AprintfromWaterhouse(1841)thatis clearlyPetauroides publish a description of a supposed volans because of the length of the tail and the hairy ears. This new species, which he named image was also reproduced in Waterhouse (1843) and Lydekker Petaurides cinereus. This was based on two specimens obtained fi-om (1896). the Bellinden-Ker Range, north- Major T.L. Mitchell collected a presumed new east Qld. The name Petaurides is a species of glider "from the banks of the Murray", definitemisspellingofPetauroidesThomas 1888 (see named it Petaurvs lencogaster and "deposited [it] Ramsay 1890). It is also noted that these specimens in the Australian Museum (AM)" (Beimett 1837; hadearlierbeenexhibitedatameetingofTheLinnean Mitchell 1838). Gray (1841) suggested that it Society ofNSWunder the name ofBelideus cinereus "may only be a variety of [the] P. taguanoides" of (Anon 1890). Waterhouse (=the greater glider) (see also mention The next taxonomic contribution was by Iredale ofP leucogaster in Gray 1842, 1843; Krefft 1864). and Troughton (1934). They argued that the generic Several authors considered leucogaster to be name Schoinobates Lesson 1842 had been published synonymous with P. volans (Gould 1863; Thomas before Petauroides Thomas 1888, and advanced the name S. volans with four subspecies: S. v. volans; Proc. Linn. Soc. N.S.W., 129, 2008 43 EARLY HISTORY OF THE GREATER GLIDER S. V. incanus; S. v. armiUatus and S. v. minor. LeSouefandBurrell 1926; Fleay 1933), 'thebrill'(De Subsequently,S. volanswas inuseforaround50years Vis 1886), 'flying phalanger' (Haswell 1886), 'great (Fleay 1947, 1968; Tate 1945;Anon 1946; Troughton flying oposssum', 'flying squirrel' (Lucas 1890), 1935, 1941; Marlow 1958, 1962; de Beaufort 1966; 'dusky glider' (Fleay 1933; Ride 1970), 'greater Ride 1970; Strahan 1980, 1981). However, the glider-possum' (Iredale and Troughton 1934; Anon nomenclatural change by Iredale and Troughton 1946), and 'greater glider' (Marlow 1958). Stability (1934) was groundless. McKay (1982) pointed out in the vernacular name was achieved in 1980 when that Schoinobates was first used by Lesson (1842) a committee of the Australian Mammal Society to supersede Petaurista leucogenys Temminck, 1838 formalised it as the 'greater glider' (Strahan 1980). {=Pteromys leucogenys; the Japanese flying squirrel). In fact, this was an error on Lesson's part because MORPHOLOGY there are no marsupials in Japan (Palmer 1904). Nevertheless, it was highly irregular for Iredale and Troughtonto amendthetype locality oiP. leucogenys The morphology was first described by Phillip from "Japan" to "Sydney". Probably, Iredale and (1789). He statedthatthe"tipofthenosetorootoftail Troughton (1934) did not view the original account [was] 20inches [=508mm],tail22 inches [=559mm], and illustration ofP. leucogenys in Fauna Japonica loins 16 inches [=406 mm]." The ears were described (Temminck 1838), which clearly depicts a sciurid. as "large and erect", the fur "glossy black" on top, Schoinobates Lesson, 1842, is therefore properly "mixed with grey", and "the under parts...white". It placed as ajunior synonym oiPetaurista Link, 1795. was noted that the fiir "continued to the claws", and Thus, McKay's (1982) assessment that the name that the membrane "expanded on each side of the Schoinobates was unavailable and that Petauroides body". Phillip (1789) also described and illustrated must stand wasjustified. the foot (Figure 1). He observed that the "fore legs Iredale and Troughton (1934) also nominated have five toes on each foot, with a claw on each; the Petaurus maximus as a synonym for the subject hinder ones four toes, with claws, (the three outside species, listingPartington(1837)astheauthority.This ones without any separation) and a thumb without a was acceptedby McKay (1982) and Flarmery (1994). claw". Following Phillip (1789), similar descriptions However, McKay later attempted unsuccessfully were also published by subsequent authors based on to track down the original reference and stated that his original accountandfromthe illustrationprovided the relevant page in the book he examined "contains (i.e. Kerr 1792; Meyer 1793; Bechstein 1800). Shaw no reference to this or any other mammal" (McKay (1794)providedmorphological descriptions basedon 1988). We note that McKay (1988) misread Iredale the illustration reproduced in Figure 2. andTroughton's (1934)referencetoPartington(1837: One diagnostic feature ofP. volans is the flying 424) because P. maximus is indeed described in The membrane that runs from the elbow to the knee, and British Cyclopaedia ofNaturalHistory, but not in the this wasnotedby several earlyzoologists (Kerr 1792; The British Cyclopedia ofArts andSciences, which Turton 1806; Desmarest 1817; Waterhouse 1841, was read by McKay (1988). After reading Partington 1846). Thomas (1888) added that the membrane is (1837) with its reference to some "almost white" "verynarrowalongthe sides ofthe forearmandlower specimens,weaccept/^ maximusas synonymouswith leg". Ramsay (1890) stated that the "parachute" or the greater glider (following Iredale and Troughton "wing membrane" commences a little in front ofthe 1934). The preceding literature review oftaxonomy elbow-joint, and extends to about half-waybelow the ofthe Greater Glider is presented in Table 1. knee-joint. Numerous early authors also noted the Common names for the subject species syndactylous hind feet (Kerr 1792; Shaw 1794, 1800; have included 'black flying opossum' (Phillip Bechstein 1800; Lacepede 1801; Tiedemann 1808; 1789), 'flying opossum' (Kerr 1792; Turton 1806; Desmarest 1820; Partington 1837). Waterhouse 1841), 'long-tailed opossum' (Shaw Some authors have compared the size of this 1794, 1800; Turton 1806; Waterhouse 1841), 'large- species to animals known from Europe. For example, tailed Petaurista', 'Peron's Petaurista' (Cuvier 1827), it has been suggested to be about the size ofa "black 'white-bellied flying squirrel' (Bennett 1837), 'grey rat" (Shaw 1800), "flying squirrel" (Desmarest flying squirrel' (Bennett 1837; Waterhouse 1841) 1803; Tiedemann 1808), "surmulot" (Cuvier 1817; 'large-tailed flying squirrel' (Bennett 1837), 'taguan Desmarest 1820; Lesson 1827), "squirrel ofEurope" flying opossum' (Waterhouse 1838b), 'taguan flying (Desmarest 1820; Lesson 1827), and "brown phalanger' (Waterhouse 1846; Thomas 1888, 1923; rat' (Partington 1837). More recently, it has been Fleay 1933), 'greater flying phalanger' (Gould 1863; suggested to be about the size of a domestic cat 44 Proc. Linn. Soc. N.S.W., 129, 2008 1 K.S. MALONEYAND J.M. HARRIS Table 1: New synonymy based on the current review. Petauroides Thomas, 1888 Petaurus Shaw, 1791 Didelphis Ken, 1792 Voluccella Bschstein, 1800 Phalanger Lacepede, 1801 P/7fl/o«g/5to Desmarest, 1803 Petowmto Desmarest, 1820 Petauroides Thomas, 1888 Petaurides Ramsay, 1890 BelideusAnon, 1890 Schoinobates Iredale and Troughton, 1934 Petauroides volans (Kerr, 1792) Petauroides volans volans (Kerr, 1792) Didelphis volans Kerr, 1792 Didelphis voluccella Meyer, 1793 Didelphis macroura Shaw, 794 1 Voluccella nigra Bechstein, 1800 Voluccella macroura 'BechsXem, 1800 Phalangervolans Lacepede, 801 1 Phalangista volans Desmarest, 803 1 Petaurus macroura Oken, 1816 Petaurus nigerOken, 1816 Petaurus taguanoides Desmarest, 1817 Petaurus macrourus Desmarest, 1817 Petaurusperonii Desmarest, 1817 Petaurista taguanoides Desmarest, 1820 Petaurista macroura Desmarest, 1820 Petauristaperonii Desmarest, 1820 Phalangista macroura Schinz, 182 Petaurus didelphoides Cuvier, 825 1 Petaurus maximus Partington, 1837 Petaurus volans Thomas 1879 Petowrato vo/a«5 Thomas 1885 Petauroides volans typicus Thomas, 1888 Petauroides volans incanus Thomas, 1923 Petauroides volans armillatus Thomas, 1923 Schoinobates volans volans Iredale and Troughton 1934 Schoinobates volans incanus Iredale andTroughton 1934 Schoinobates volans armillatus Iredale and Troughton 1934 Petauroides volans minor (Collett, 1887) Petaurista volans minor Collett, 1887 Belideus cinereus Anon, 1890 Petaurides cinereus Ramsay, 1890 Schoinobates volans minor Iredale and Troughton 1934 (Flannery 1994). Le Souefand Burrell 1926). Gould (1863) stated that Colouration was also frequently commented on. "it is subject to very great variety in the colouring of Forexample,Cuvier(1817)reportedthatthefurexists its fiir, some specimensbeing entirelyblackishbrown in different tones ofbrown, with many varieties, and [see Figure 4], grey to cream and others quite white", others are whitish. Fully white specimens were also Kreflft (1871) reported that the species "varies much noted (Lesson 1827; Waterhouse 1841; Krefft 1864; from creamy-white to spotted black and white and Proc. Linn. Soc. N.S.W., 129, 2008 45 EARLY HISTORY OF THE GREATER GLIDER Figure 4: Petaurista taguanoides from Gould (1863) (=P. volans). Note: The front arms ofthe back- ground glider are shown in the wrong position as P. volans tucks them under the chin when gliding (Fleay 1933; Grzimek 1967; McKay 1989). 46 Proc. Linn. Soc. N.S.W., 129, 2008 K.S. MALONEYAND J.M. HARRIS Table 2: Dental formulas provided in the early natural history literature forPetauroides volans. Abbre- M viations: I = Incisors; C = Canines; = Molars; P = Premolars. For Thomas (1885, 1888) an asterisk indicates that the tooth is sometimes or commonly absent. Source Dental formula Desmarest 1820 6^11 11 66 77 /,-;C, or ;M or = 32or34 2 00 22 66 66 Cuvier 1825, 1826 6 00 8 8 I—,C, ;M, = 38 2 00 68 — Lesson 1827, 1830, 1838; 6 00 8 8 Fischer 1829 /-;C, ;M, = 38 2 7 7 — Waterhouse 1838b 33 11 33 44 / ;C, ,P--\M, = 34 11 00 11 44 Krefft 1871; Collett 1887 —33 —11 —33 44 / ;C, ;P ;M, = 40 11 11 3 3 4 4 Thomas 1885 / 12 3 C—1 P 12 3 M 1234 x2 = 34or40 10 1* 1*2*3 12 3 4 Thomas 1888 123 — 034 1234 / C, -1P, 1 {or\) M, = 17 + (atmost) 3 (or 16 + 4) x 2 =40 12*0 1* 3*4 123 4 perfect black, beneath the fiir is always white." Le DENTITION Souef and Burrell (1926) stated that "as a rule [the] colour [is] darker in winter than in summer." They Phillip (1789) stated that in "the upper jaw also stated that "animals from Gippsland (Victoria) forwards are four small cuttingteeth, thentwo canine [were] dead black above and on tail; pure white on ones, and backwards five grinders: the under jaw NSW undersides", whereas Qld and specimens were has two long large cutting teeth, five grinders, with "usually smoky grey" and "white specimens [were no intermediate canine ones, the space being quite reportedly] common." vacant". Similarlywordeddescriptionswereprovided Other notable morphological features described by Kerr (1792) and Turton (1806). A in the early literature include the ears, tail and size dentalformulaforthespecieswasfirstprovided differences between the sexes. Waterhouse (1838a, by Desmarest (1820) (see Table 2). He counted six 1841, 1846) stated that "the ears are entirely covered upperandtwo lowerincisors,butwasuncertainabout externallywith longanddense fur, flesh-colouredand thenumberofcaninesandpremolars.Thisuncertainty almost bare within" (see also Krefft 1864, Thomas led him to indicate a total of 32 or 34 teeth. Cuvier 1888, Ramsay 1890). The tail was reported as not (1825) and Lesson (1827) counted a total of38 teeth. being prehensile (Lacepede 1801; Tiedemann 1808; Cuvier (1825) reported that the space between the Partington 1837), and longer than the body (Shaw incisors and molars is occupied by two rudimentary 1800; Turton 1806; Cuvier 1817). Thomas (1888) teeth. Waterhouse (1838b, 1841) and Owen (1841, described and illustrated the naked tip of the tail. 1845) mentioned they had never observed any of Gould (1863) stated the "sexes offer no external these diminutive teeth in the specimens they had difference, except that the female is somewhat examined. Waterhouse (1841) suggested that Cuvier smaller than the male" (see Flarmery 1994, as this is (1825)mayhaveinadvertentlydescribedthedentition erroneous). Various other aspects ofthe morphology of Phalangista cookii {=Pseudocheirus peregrinus; of this species are discussed in the literature, but common ringtail possum). These two species do lack of space precludes a detailed discussion here. have great similarity in their dental characteristics, as However, these aspects include skull structure noted by early zoologists (Owen 1841, 1845; Giebel (Waterhouse 1846; Collett 1887; Thomas 1888) and 1853, 1855; Thomas 1885; Collett 1887) and more myology (Haswell 1886). modemauthors (Tate 1945; Triggs 1996). Waterhouse (1838b) provided a dental formula indicating a total of 34 teeth. Subsequent authors concurred with this Proc. Linn. Soc. N.S.W., 129, 2008 47 EARLY HISTORY OF THE GREATER GLIDER observation (Waterhouse 1841, 1846; Wagner 1843; flowers, andpreferring thebarkofthebranchestothe CoUett 1887; Ramsay 1890). Early illustrations ofthe leaves" (Le Souef and Burrell 1926). Fleay (1933) dentition in Cuvier (1825, 1827), Waterhouse (1846) stated "one of the chief difficulties in captivity is and Giebel (1853, 1855) support the dental formula the maintenance ofan abundant supply ofthe tender ofWaterhouse (1838b). leavesofacceptablespeciesofeucalypts"andreported Krefft's (1871) dental formula (Table 2) was for on collecting trips to obtain sufficient amounts of a total of 40 teeth (see also Collett 1887). Thomas' leaffrom E. elaeophora and E. australiana. He also (1885) assessment was that the number of teeth reported that "captive specimens could be persuaded varied from 34 to 40, dependent on the presence or toacquireanadditionaltasteforbreadandmilkspread absence of a small canine and two premolars in the with a sweet jam, but only as an adjunct to the diet lower jaw. Thomas (1888) attempted to improve his ofeucalypt leaves." Grzimek (1967) stated "because earlier dental formula by changing the position of \P. volans'] are exclusive in their diet, like koalas, no the lower canine to the incisor position (Table 2), specimen has ever reached a European zoo alive." and remarked that the "presence or absence of the Menkhorst and Knight (2004) stated that it "eats minute teeth is not of any systematic importance". only eucalypt leaves and buds." However, Maloney Thomas (1888) provided illustrations of the upper and Harris (2006) report feeding observations from andlowerjaw ofP. v. volans andP. v. minor, although several non-eucalypts. these are notconsistentwithhis dental formula. Later Interms ofhabitat, Gould (1863) wrotethatit"is reviewers have alluded to a socket in the lower jaw strictly an inhabitant ofthe extensive brushes which where a small incisor would be present (i.e. Archer stretch alongthe south-eastern andeasternportions of 1984; Triggs 1996). Twenty-one P. volans specimens New SouthWales". It has also been reported to occur AM in the were recently examined by us, and four in Eucalyptus iorQsis (Le Souef and Burrell 1926, (19%) were noted to have minute teeth between the Anon 1946). Fleay (1933) stated that the species was incisors andpre-molars. found"favouringthe tallertimberareas andgenerally inhabiting dead trees in the gullies of mountainous country". Marlow (1958) reported thatP. volans was HABITATAND DIET more abundant in dry than wet sclerophyll forests and less common in open woodland. Ride (1970) Some information on the habitat and diet of stated that "the habitat is sclerophyll forest and tall P. volans is available in the early literature. Gould woodland". (1863) stated that the species seeks "blossoms ofthe Eucalypti...together with the tender buds and shoots of the same trees". Similarly, Le Souef and Burrell DISTRIBUTIONANDABUNDANCE (1926) statedthatthe "food consists ofthe leaves and buds ofeucalyptus-trees". They also added that: The earliest statements on the distribution ofthe NSW subjectspecieswasthatitinhabits (Phillip 1789; 'careful examination ofthe contents of Kerr 1792) or "New Holland" (=Australia) (Meyer several stomachs ofanimalstakenfrom 1793; Shaw 1794; Cuvier 1798; Bechstein 1800). The the forests has not revealed anything earliest specific localities mentioned were for places else, but in the Myall Lakes district inNSW, i.e. Botany Bay, Port Jackson, Sydney, Blue PSTSW]... we have observed this Mountains, Port Macquarie, Bathurst, Maitland, species on the casuarina-trees; in one Clarence River and Goulbum Plains (Cuvier 1826, such case the contents ofthe stomach, 1827; Lesson 1830; Bennett 1837; Waterhouse 1841; although much masticated, seemed to Gray 1841; Krefft 1864). Other early distributional NSW AM be the casuarina-leaves. Mr. Ralph C. records for include Sutherland (1908, AM Blackett, forest ranger at Queanbeyan M2003), Helensburgh (1909, M2051), Bowral AM AM [NSW]..., states that they chiefly (1918, M2724), Myall Lakes (1922, feed on E. regnans, and to a lesser M33762), Gerringong and Milton (Troughton 1935, extent on E. viminalis, E.fastigata, E. 1941), Geehi Gorge (Mt Kosciuszko area) (Anon australasiana, andothernarrow-leaved 1946), Armidale and Tidbinbilla Nature Reserve peppermints.' (1974), (see Maloney and Harris 2006). Early literature records from Queensland are In captivity, P. volans has been observed to eat north of the Herbert River (de Vis 1886), Herbert E. sieberiana readily, "being especially fond of the Vale, Coomooboolaroo, Calliungal (Collett 1887), 48 Proc. Linn. Soc. N.S.W., 129, 2008